SHOREBIRD MIGRATION AT NORMAN, OKLAHOMA: 1961-63

LEWIS W. ORING AND W. MARVIN DAVIS

EPEATED censusing of a limited area provides a valuable source of quanti- R tative migration data despite the many variables involved. Though several such studies of shorebirds have been conducted on or near the Atlantic and Pacific coasts (Urner, 1929; Urner and Storer, 1949; Storer, 1951; Dyke, 1955) no comparable data are available from the central flyway region. This report is a result of observations made from 1 to 15 No- vember 1963 at a small impoundment in Norman, Oklahoma.

DESCRIPTION OF AREA

Norman, Oklahoma is adjacent to the floodplain of the Canadian River. The river flows generally from west to east across the state, but follows a northwest to southeast direction for a considerable distance on both sides of Norman. Native woody vegetation near the city is limited to areas along the river and other streams. Max Westheimer Airfield was developed as a military base in 1942, and is presently operated by the University of Oklahoma. More than 200 acres of asphalt remain, considerable portions of which are not used as runways or taxiways. One such unused area has been the site of a pond since 1949 or 1950. At that time a drainage ditch was dammed in the pasture adjacent to the west edge of the airfield. Except for periods of extreme drought, the pond extends from the pasture onto the airfield as a shallow pool having a maximum depth of about two feet, a length of about 300 yards, a width of about 70 yards, and an area of about three acres. Maximum size is not regu- larly maintained, however, as an extremely high evaporation rate prevails in Oklahoma from April to October. During these months of 1961-63 the average dimensions of the pond were 150 x 40 yards and the average surface was 1.4 acres. The broad, shallow portion of the impoundment, in contrast to the deeper and narrower section in the pasture, has proved quite attractive to migrant shorebirds despite its small size and lack of sandy or muddy shores. When first laid, the asphalt that underlies and nearly surrounds the pond constituted a primary bare area. This area has been invaded for some years by a vigorous plant growth distributed linearly along the seams and cracks of unused areas. Such growth invades the pond at a few points. For about 100 yards along the southwest shore of the pond the asphalt was broken up and hauled away several years prior to this study. This section constitutes the

166 Oring and SHOREBIRD MIGRATION Davis 167

major site of both submerged and emergent vegetation and is the only area of the pond in which an automobile cannot be driven.

COVERAGE

Censuses were usually taken at least once daily from 1 March to 31 October of all three years. At times, however, a day was missed and on three occasions (1-8 ; 25-30 ; and 26 August to 3 ) both observers were absent from the study area for about a week. From 1 March 1961 to 31 daily visits were made at 0600 and lSO0 CST. Throughout the remainder of the study the time of day at which observations were made was variable. Observation periods averaged about an hour, but some lasted from three to six hours in May and September.

INFLUENCE OF WEATHER

During the frequent spring periods of clear skies and strong southerly winds we experienced a rather uniform pattern of migration and found few rare species, whereas at times of heavy cloud cover, fog, or northerly winds, migrants were grounded and we regularly found uncommon species and larger numbers of individuals. It seems likely that a relatively constant flow of shorebirds passes through central Oklahoma during the height of the north- bound migration but that many of these birds are seen only when forced to the ground by inclement weather. Occasionally an extended wet spell occurred in spring. Concurrently there was a reduction in the shorebird population due apparently to two factors: (1) an increase in feeding sites caused by accumulation of water in fields and on the runway, and (2) the reduction of available food at the pond due to the elevated water level. During very heavy rain or hail only the largest shorebirds remained at the pond-others went to less exposed areas away from the water. Shelter was also sought when the wind velocity reached about 30 mph. At these times we found large flocks of many species running about far from the water feeding voraciously on grounded insects and seeds. Summer in Oklahoma is characterized by an almost continuous span of variably intense southerly winds and clear skies. During such periods the number of southbound migrants changed little from day to day. But when the first cold snap arrived with stron g northerly winds, usually sometime in September, a great variety of shorebirds suddenly appeared; and this phe- nomenon was repeated to a lesser degree with each ensuing north wind until the supply of migrants was exhausted sometime in November. Our observa- tions of fall migrants are thus consistent with the findings of Brooks (1965) THE WILSON BULLETIN 168 Vol. 78, No. 2 for shorebirds and Hassler et al. (1963) f or p asserines in that they considered the shift of wind to the north to be the primary weather stimulus for south- bound migration. The paucity of migrants during the days just after an extended stormy period was also notable. In sprin g this may have been due to the reluctance of birds to leave the Gulf Coast or other areas of concentration during times of weather disturbance. In summer and fall it may have been due to the small number of birds physiologically ready to migrate that still remained at northern “staging points” following a storm front.

RESULTS

Thirty of the 39 species of shorebirds recorded from Oklahoma were seen in the study area. Of the nine unrecorded species (Piping Plover, Charudrius melodus; Mountain Plover, Eupoda montana; Ruddy Turnstone, Arenaria interpres; Woodcock, Philohela minor; Eskimo Curlew, Numenius borealis; Knot, Calidris canzztus; Short-billed Dowitcher, Limnodromus griseus; Black- necked Stilt, Himantopus mexicanus ; and Red Phalarope, Phalaropus fuli- carius), the Knot and Red Phalarope have never been collected in Oklahoma and only the turnstone is of more than accidental occurrence in the central part of the state. The Whimbrel (Numenius phaeopus) , White-rumped Sandpiper (Erolia fuscicollis) , Marbled Godwit (Limosa fedoa) , Hudsonian Godwit (L. hae- mastica), Sanderling (Crocethia alba) , and Northern Phalarope (Lobipes Zobatus) were recorded from only the northward migration whereas the Snowy Plover (Charadrius alexandrinus) , Dunlin (Erolia alpina) , and American Avocet (Recurvirostra americana) were seen only during the south- bound season. Despite increased populations following the reproductive season, the Golden Plover (Plzzvialis dominica), Spotted Sandpiper (Actitis macularia) , Greater Yellowlegs (Totunus melunoleucus) , Pectoral Sandpiper (Erolia melunotos) , Bairds’ Sandpiper (E. bairdii) , Buff-breasted Sandpiper (Tryngites subruficoZZis), and Wilsons’ Phalarope (Steganopus tricolor) were all much less common during southward than northward migration. Our coverage of the study area and its apparent attractiveness to shorebirds were relatively constant throughout the year and thus cannot account for this paradox. Conversely, the Western Sandpiper (Ereunetes mauri) was numer- ous during the southward migration, but recorded on only four dates in spring. The Upland Plover (Rartramia longicaudu) , Solitary Sandpiper (Tringa solitariu) , Least Sandpiper (Erolia minutiZZu), Long-billed Dowitcher (Limnodromus scolopaceus), and Stilt Sandpiper (Micropalama himantopus) were also much more common in fall than in spring. Among these, only the Solitary and Western sandpipers showed an increase in numbers greater than SHOREBIRD MIGRATION 169

TABLE 1 RANKING OF SHOREBIRDS BY ABUNDANCE IN NORTHWARD MIGRATION, 1961-63

Maximum Species Rank Total Total dates individuals daily totals Bairds’ Sandpiper 1 1% 2,553 150 White-rumped Sandpiper 2 86 1,524 105 Semipalmated Sandpiper 3 76 900 145 Buff-breasted Sandpiper 4 46 1,027 215 Wilsons’ Phalarope 5 56 1,071 140 Pectoral Sandpiper 6 117 857 75 Least Sandpiper 7 73 877 75 Lesser Yellowlegs 8 104 848 64 Greater Yellowlegs 9 64 587 50 Golden Plover 10 36 269 35 Stilt Sandpiper 11 31 88 17 Long-billed Dowitcher 12 31 95 11 Upland Plover 13 23 79 15 Spotted Sandpiper 14 39 68 6 Semipalmated Plover 15 14 29 7 Common Snipe 16 18 22 3 Hudsonian Godwit 17 6 14 5 Willet 18 6 13 6 Sanderling 19 6 13 4 Black-bellied Plover 20 2 13 7 Marbled Godwit 21 4 8 3 Western Sandpiper 22 4 5 2 Solitary Sandpiper 23 5 5 1 Long-billed Curlew 25 1 1 1 Whimbrel 25 1 1 1 Northern Phalarope 25 1 1 1 that explainable by the addition of offspring to the population. The Semi- palmated Plover (Charadrius semipalmatus) , Killdeer (C. vociferus) , Com- mon Snipe (Capella g&&ago), Lesser Yellowlegs (Totanus fluvipes) , and Semipalmated Sandpiper (Ereunetes pusillw) were about as common during the northward as during the southward migration. The relative abundances (after Urner and Storer, 1949) of 26 species of shorebirds found in northward migration, and of 23 species seen in southward migration are presented in Tables 1 and 2. The Killdeer is excluded from this comparison because of its year-round presence. Seasonal extreme dates for the three-year span are given below for each species. Migration peaks and dates of maximum counts for both northward and southward seasons are given where data are adequate. It must be remembered that our area does not provide optimum habitat for all species of shorebirds. Common Snipe and Solitary Sandpipers are more June 1966 170 THE WILSON BULLETIN Vul. 78, No. 2

TABLE 2 RANKING OF SHOREBIRDS BY ABUNDANCE IN SOUTHWARD MIGRATION, 196163

Total Maximum Species Rank individuals daily totals Least Sandpiper 1 207 1,892 51 Lesser Yellowlegs 2 123 652 70 Western Sandpiper 3 96 429 30 Semipalmated Sandpiper 4 106 336 23 Pectoral Sandpiper 5 74 300 30 Bairds’ Sandpiper 6 69 350 21 Buff-breasted Sandpiper 7 30 246 50 Long-billed Dowitcher 8 49 153 29 Stilt Sandpiper 9 57 174 14 Upland Plover 10 49 113 11 Solitary Sandpiper 11.5 44 71 4 Wilsons’ Phalarope 11.5 16 52 10 Greater Yellowlegs 13 24 38 6 Semipalmated Plover 14 10 16 3 Common Snipe 15 18 26 3 Golden Plover 16 15 21 4 Dunlin 17 4 14 7 American Avocet 18 3 9 7 Spotted Sandpiper 19 9 9 1 Black-bellied Plover 20 6 7 2 Willet 21.5 7 7 1 Long-billed Curlew 21.5 2 5 4 Snowy Plover 23 1 1 1

common along marshy lake edges and streams; Upland Plovers are abundant in alfalfa and plowed fields; and Snowy Plovers are regular along sandy riverbeds. All of the larger forms-Marbled Godwit, Hudsonian Godwit, Long-billed Curlew (Numenius americanus), Whimbrel, Willet (Catoptro- phorus semipalmatus), and American Avocet, as well as Black-bellied Plover (Squaturola squatarola), Dunlin, and Sanderling-are more regular at the larger reservoirs. This may be due to the differences in sample sizes. For the remaining species the study pond appears quite suitable and for some, e.g., Buff-breasted Sandpiper, superior. In the height of spring migration it was obvious that most flocks were restless and little inclined to linger more than a few hours. According to our observations, few birds other than Bairds’ Sandpipers and Upland Plovers remained longer than a day. During most of the southward migration, how- ever, migrants faced a head wind and under such conditions often lingered several days. Determination of which birds were new to the census and which had al- Oring and SHOREBIRD MIGRATION Davis 171 ready been counted was complicated by the fact that certain birds circulated back and forth between the study area and another nearby pond. In this re- gard, we depended upon flock size, direction of departing and arriving groups, and presence of crippled, banded, or otherwise marked individuals.

SPECIES STATUS

Semipalmated Plover.-Extremes: 17 April6 May and 26 July-25 September. One recorded 11 . Maximum counts: 25 (7) and 22 April 1963 (5). Most common at pond but also noted on runway far from water. Snowy Plover.-Observed only once, 19 ; despite its breeding three miles south of the study area at the Canadian River. Migrants were often seen at river in April and May. Killdeer.-Nests commonly, winters sparingly. Peak migration: 2-18 March and 5 August-l November. Maximum counts: 14 March 1961 (u)), 16 March 1961 (111, 16 (8), 29 (125), 5 August 1961 (451, and 20 August 1961 (40). Seemingly migratory flocks in summer were noticeably larger during midday. One adult banded 13 remained at least until 8 . Golden Plover.-Extremes: 26 March-22 May and 20 July-11 November. Peak mi- gration: l&27 April. Maximum counts: 15 April 1961 (41), 10 April 1961 (400)) 16 April 1961 (351, and 8 September 1963 (4). In fall larger numbers were seen in plowed fields three miles northwest of study area. In spring it was most common in evening and early morning. They favor a small section of pond shore that has scattered chunks of broken asphalt; also frequent nearby fields and runway. Only two of 269 in spring were in complete breeding plumage-one 18 May 1961, the other 21 May 1961. Black-bellied Plover.-Extremes: 18 and 19 May; and 2 August-28 October. Maximum counts: 18 May 1961 (7) and 19 May 1961 (6). Both spring flocks were seen at day- break; all birds were in full breeding plumage. Common Snipe.-Extremes: 26 March-28 April and 23 September-31 October. Peak migration: 3-20 April and 23 September-31 October. Winters near Norman in favor- able habitats. Long-billed Curlew.-Single birds were observed 25 April and 4 ; flock of four 19 July 1963. Whimbrel.-A single bird was observed 1 June l%l. Upland Plover.-Extremes: 11 April-18 May and 6 July-18 October. Four mid-June records of single birds indicate possible local nesting. Peak migration: 12230 April and 2 August-24 September. Maximum counts: 12 April 1963 (151, 30 (71, 24 and 28 April 1961 (6), 9 September 1962 (ll), 3 September 1961 (8), 3 August and 7 September 1963 (7). First southward movement, as determined by nocturnal flight calls, 6 July. Usually seen in fields near pond, but occasionally seen feeding or bathing at water. Much more numerous in alfalfa and plowed fields west of Norman. Spotted Sandpiper.-Extremes: 19 April-31 May and 12 July-12 September. Peak migration: 27 April-23 May. Maximum count: 13 (6). Only singles ob- served in summer. Seen several times resting on the asphalt far from water. Solitary Sandpiper.-Extremes: 9 April-24 May and 6 July-25 August. Maximum count: 24 (4). Willet.-Extremes: 27 April6 May and 30 June-4 September. Maximum count: 27 April 1963 (6). Summer observations were all of single birds. Greater Yellowlegs.-Extremes: 8 March-7 May and 9 July-12 October. Peak mi- THE WILSON BULLETIN June 1966 172 Vol. 78, No. 2 gration: 25 March-20 April and 15 July-9 September. Maximum counts: 9 April 1962 (50), 6 April 1962 (49), 7 April 1962 (42), 9 September 1962 (6), 20 (4), and 26 (3). Most common during April, in unmixed flocks arriving at or after sunset. In contrast to Lesser Yellowlegs it was never seen away from water. Lesser Yellowlegs.-Extremes: 7 March-3 June and 21 June-28 October. Peak mi- gration: 29 March-12 May and 26 July-6 September. Maximum counts: 24 April 1963 (64), 28 April 1963 (63), 25 April 1963 (41), 3 September 1961 (70), 4 September 1961 (55), and 9 September 1962 (28). Usually observed wading in water but frequently seen in sheltered areas far from water during storms or strong winds. Pectoral Sandpiper.-Extremes: 2 March-3 June and 9 July-17 October. Peak mi- gration: 10 April26 May and 26 Julyd September. Maximum counts: 4 May 1961 (75), 30 April 1963 (36), 27 March 1961 (28), 3 September 1961 (31), 5 September 1961 (30), and 25 August 1963 (16). A single bird, apparently male, seen 23 June 1962 was presumed to be an early southward migrant. Usually they favor the pond shore although large numbers land on the runway or in the adjacent pasture. After rain they often visit puddles along the runway. The sexes rarely migrate together in spring. In 1961, Dan R. Harlow collected 124 Oklahoma specimens for parasitological analysis (81 in spring; 43 in summer and fall). His data are as follows: to 5 April-27 males, 0 females; 9 April-11 males, 1 female; 11 April to 25 April--S females, 4 males; and 27 April to 2 June-33 females, 0 males. Extreme dates for sexes during spring mi- gration in study area (as verified by specimens) are as follows: males 16 March to 6 May; females 9 April to 3 June. Data from 50 fall Oklahoma specimens (including Harlows’ 43) yield the following date extremes by sex: males 25 July to 26 October; females 31 July to 26 October. All of Harlows’ fall specimens were osteologically immature. This, coupled with the decrease in total Pectorals observed in southward migration, suggests that part or all of the adult population follows a different southward path and that the flocks found here durin, w summer and fall contain juvenile birds of both sexes. White-rumped Sandpiper.-Extremes: 27 April-21 June. No fall records. Peak migration: 3-26 May. Maximum counts: 16 May 1963 (105), 20 May 1963 (65), and 20 May 1962 (54). Most common at midday. Frequents pond shore most often but also regular on the runway and in the plowed fields. Sexes tend to migrate separately. In Oklahoma, males collected from 2 May to 26 May; females from 17 May to 6 June (data gathered by University of Oklahoma Museum of Zoology). Bairds’ Sandpiper.-Extremes: 3 March-31 May and 20 July-13 October. Peak Mi- gration: 12 April-24 May and 20 July-24 September. Maximum counts: 10 May I963 (150), 9 May 1963 (145)) 17 May 1963 (135), 4 September l%l (21), 3 September 1961 (20)) and 25 July 1962 (20). This species often remains several days and in spring some may remain a week or longer. Often seen on runway or in adjacent fields, especially when standing water is widespread or when winds are exceptionally strong. Least Sandpiper.-Extremes: 27 March-24 May and 4 July-l November. Peak mi- gration: 22 April-18 May and 8 July-22 October. Maximum counts: 1 May 1961 (75), 8 May 1961 (65), 8 May 1963 (58), 6 (51), 4 September 1961 (50), and 8 October 1961 (41). Most common during midday. Usually found at ponds’ edge but seek sheltered areas in heavy winds. Dunlin.-Extremes: 9-29 October. Four records only: 9 October 1961 (7)) 10 October 1961 (5), 11 October 1961 (l), and 29 October 1961 (1). Long-billed Dowitcher.-Extremes: 16 March-18 May and 10 July-20 October. Peak migration: 6 April-8 May and 26 July-18 October. Maximum counts: 29 April 1962 Oring and Dark SHOREBIRDMIGRATION 173

(II), 27 April 1962 (71, 22 April 1963 (61, 6 October 1961 (29)) 18 October 1961 (II), and 23 September 1961 (IO). On a few occasions the presence of Short-billed Dowitchers was suspected Stilt Sandpiper.-Extremes: 20 April-25 May and 6 July--25 September. Peak mi- gration: 25 April-16 May and 2 August-25 September. Maximum counts: 10 May 1963 (6)) 9 May 1963 (6), 11 May 1962 (5), 5 September 1962 (141, 3 September 1961 (IO), and 12 August 1961 (IO). Semipalmated Sandpiper.-Extremes: 8 April-30 May and 24 June-13 October. Singles observed 11 and 16 . Peak migration: 18 April-26 May and 12 July- 3 September. Maximum counts: 25 April 1961 (1455), 26 May 1961 (681, 4 May 1962 (571, 30 August I%1 (23), 1 September 1961 (181, and 3 September 1961 (15). Most common at midday. Occasionally seen on the runway and in plowed fields. Western Sandpiper.-Extremes: 16 April-27 April and 30 June-9 October. Peak mi- gration: 2 July-3 September. Maximum counts: 29 August 1961 (301, 27 August 1961 (25)) and 17 July 1963 (20). Buff-breasted Sandpiper.-Extremes: 17 April-3 June and 23 July-4 October. Peak migration: 4-19 May and 9 August-8 September. Maximum counts: 9 May 1962 (2151, 10 May 1962 (117), 12 May 196.2 (98), 9 August 1963 (431, 5 September 1962 (291, and 8 September I%3 (26). Most common at midday. They feed mainly in nearby fields and on the runway, but bathe and display at the pond. Flocks in April and early May contained a preponderance of males while those after the middle of May often were composed solely of females. Marbled Godwit.-Extremes: 26 March-l May. Four records only: 26 March I%1 (31, 9 April I%1 (l), 24 April 1963 (3), and 1 May 1963 (I). Hudsonian God&-Extremes: 15 April-20 May. Maximum counts: 15 April 1961 (5)) and 17 May 1%3 (4). All birds were in breeding plumage Sanderling.-Extremes: 7-26 May. Maximum counts: 14 May 1961 (41, 11 May 1961 (3), and 8 May 1%1 (3). No fall records despite their being regular at a large reser- voir 30 miles north of Norman. American Avocet.-Extremes: 5 July-4 October. Four records only: 5 July 1962 (1)) 29 July 1%1 (71, 24 September I%1 (I), and 4 October 1961 (I). No spring records despite several occurrences near Norman. Nine of ten southbound birds lacked breeding dress. Wilsons’ Phalarope.- Extremes: 10 April-28 May and 16 July-24 September. Peak migration: 22 April-17 May and 12 August-4 September. Maximum counts: 28 April 1962 (140), 22 April 1963 (1251, 29 April 1962 (Ill), 13 August 1961 (lo), 12 August 1961 (8), and 4 September I%1 and 1962 (8). Most numerous at midday. In spring, very common for only a few days. Flocks usually are seen at the ponds’ edge, but sometimes they swim in deep water and, at times, feed on dry land. Both sexes migrate together but flocks during April contain a great preponderance of females, while those in early May often contain pairs, and flocks in mid- and late May contain more males than females. Northern Phalarope.-Single female in breeding plumage seen on 17 May 1963.

SUMMARY

A study of shorebird migration was conducted at Norman, Oklahoma, from 1 March I%1 to 15 . A total of 30 species was observed including 27 during the northward and 24 in southward migration. Bairds’ Sandpiper was by far the most abun- June1966 174 THE WILSON BULLETIN Vol. 78, No. 2 dam species during the northward migration; Least Sandpiper was the most abundant during the southbound season. White-rumped Sandpipers were common in spring but unrecorded after 21 June. Likewise, all six Sanderling records were in the spring. Species that were definitely more common in spring than in fall included the Golden Plover, Spotted Sandpiper, Greater Yellowlegs, Pectoral Sandpiper, Bairds’ Sandpiper, Buff- breasted Sandpiper, and Wilsons’ Phalarope. The Western Sandpiper was numerous dur- ing the southward migration but only five individuals were recorded during spring. The Dunlin was seen only four times, all in fall. The following species were also much more common in fall than in spring: Upland Plover, Solitary Sandpiper, Least Sandpiper, Long-billed Dowitcher, and Stilt Sandpiper. Only the Solitary and Western sandpipers, however, showed an increase in numbers greater than that explainable by the summers’ reproduction.

ACKNOWLEDGMENTS

Appreciation is expressed to Dan R. Harlow for generously allowing us access to his data on the Pectoral Sandpiper, to David F. Parmelee for critically reading the manu- script, and to Willet T. Van Velzen and John Janovy, Jr. for contributing substantially to our compilation of observations. We especially wish to thank George M. Sutton for his cooperation and encouragement throughout the study, for allowing us the use of the University of Oklahoma Museum of Zoology facilities, and for advice regarding manu- script preparation.

LITERATURE CITED

BHOOKS,W. S. 1%5 Effect of weather on autumn shorebird migration in east-central Illinois. Wil- son Bull., 77:45-54. DYKE, S. H. 1955 Shorebirds on the Conejohela flats: a five year summary. Atlantic Naturalist, 10 :260-268.

HASSLER, S. S., R. R. GRABER,AND F. C. BELLROSE 1963 Fall migration and weather: a radar study. Wilson Bull., 75:56-77. STORER,R. W. 1951 The seasonal occurrence of shorebirds on Bay Farm Island, Alameda County, California. Condor, 53:186-193. URNER, C. A. 1929 The southward shore-bird flight on the New Jersey coast in 1928. Auk, 46: 311325.

URNER, C. A., AND R. W. STORER 1949 The distribution and abundance of shorebirds on the north and central New Jersey coast, 1928-1939. Auk, 66:177-194.

DEPARTMENT OF ZOOLOGY, THE UNIVERSITY OF OKLAHOMA, NORMAN, OKLA- HOMA, 23